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Patron: Her Majesty The Queen Rothamsted Research Harpenden, Herts, AL5 2JQ Telephone: +44 (0)1582 763133 WeB: http://www.rothamsted.ac.uk/ Rothamsted Repository Download A - Papers appearing in refereed journals Clark, W. C. 1961. A revised classification of the order Enoplida (Nematoda). New Zealnd Journal of Science. pp. 123-. The output can be accessed at: https://repository.rothamsted.ac.uk/item/96yy1/a- revised-classification-of-the-order-enoplida-nematoda. © Please contact [email protected] for copyright queries. 13/11/2019 12:02 repository.rothamsted.ac.uk [email protected] Rothamsted Research is a Company Limited by Guarantee Registered Office: as above. Registered in England No. 2393175. Registered Charity No. 802038. VAT No. 197 4201 51. Founded in 1843 by John Bennet Lawes. SYSTEMATIC AND MORPHOLOGICAL STUDIES ON SOME ENOPLIDA (NEMATODA) WITH SPECIAL REFERENCE TO THE SOIL FAUNA OF NEW ZEALAND by W. C. CLARK, M.Sc. Hons. (N.Z.), D.I.C. A THESIS PRESENTED FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in the Faculty of Science of the University of London Aoril 1961 Deoartment of Nematology, Rothamsted Exoerimental Station, Harpenden, Herts. ABSTRACT This study falls in two parts, systematics and mor- phology. Systematics. The classification of the order Enoplida (Nematoda) is revised. At the subordinal level the classifi- cation is nrimarily based on the nature, number and Position of the oesophageal glands and their ducts. Five suborders are recognised; Enojlina, Alaimina, Dorylaimina, Tricho- syringina and Diectonhymatina. The new features of the classification are:- Alaimina established for the Alaimoidea and the Diphtheronhoroidea on the oesophageal and male sexual characters. The similarities between the Diphtherophoroidea and the Dorylaimoidea are considered to be due to convergent evolution. On oesophageal characters the Mononchidae are removed from the Tripyloidea and together with the Bathyodon- tidae form the Mononchoidea within the Dorylaimina. The Bathyodontidae is formed of genera previously placed in the Nygolaimidae which possess a cylindroid oesophagus, adhesive organs and a mononchoid stoma. The definitions and constitu- tions of the families Nygolaimidae and Belondiridae are modified to make these families more homogeneous. The Mermithidae are Placed with the Trichuroidea in Ward's Trichosyringina thus uniting all the families with the Oesophageal glands in the form of stichocytes. The probable course of evolution in the soil dwelling enoplids in general, and the Mononchidae in particular is described. Morphology. The monorchic male reproductive system of the Alaimidae, and the two types of oesonhago-intestinal junctions in the Mononchidae are described for the first time. New species in the genera Amphidelus, Mononchus, Iotonchus, Cobbonchus, Mylonchulus, Pungentus, Longidorus, Sectomema, Dorylaimellus and Actinolaimus are described. A new leptonchid genus is erected, and its type species described. Existing snecies of the genera Cryptonchus, Mononchus, Prionchulus, Iotonchus and Cobbonchus are re- described. CONTENTS I. Introduction Material and Methorls 1. Materials 2. Scqn-fling iochnicluo and Procedure 3. Storage of Sauulas 4. Extraction of samils 5. Fixation 6. Mounting 7. Examination 8. Measurements III. Higher Taxonomy 1. A Revised Classification of the Order Enoplida (printed) 2. The Relationships of the Soil Dwelling Eno'ilids .1V. Suborder Enoplina 1 Family Ironidae V. Suborder Alaimina 1. Description of Anrnhidolus suiralis n. sp, 2. The Male Reuroductive System in the Alaimoidea VI. Suborder Dorylaimina ; Family Mononchidae 1. Introduction 2. The Mononchidac and Biological Control 3. The Oesouhago-Intestinal JuncEion in the Mononchidae (Enoulida, Numatod-1) (printed) 4. Re-descriptions of Mononchus truncatus Bastian, M. pauillatus Bastian and Prionchulus muscorum (Duj-Irdin) (Enoulida, Norltoda). (printed) 5. The New Zealand Mononchie (Enoplida, Nematoda) I. The genera Mononchus Bastian and Prionchulus Cobb. (printed) 6. Tho Mononchidac (Enonli a, Nematoda) of New Zealand II. The genus Iotonchus (Cobb, 1916) Althorr, 1950. (printed) 7. Mononchid NomatoCa) of New Zealand Review of the genus Cobbonchus Andrassy, 1958 with descriptions of n.)w SDOCLDS. (printed) 8. The Mononchidae (Enoplia- Nimatoda) of New Zealand IV. The genus Mylonchulus (Cobb, 1916) Pennak, 1953 (printed) 9.. Evolution within the 11thonchidae VII. Superfamily Dorylaimoidea I. Dorylaiminae; Description of Pungentus maorum n.sp. 2. Actinolaiminae; Description of Actinolaimus zealandicus n. sp. 3. Tylencholaimina; Description of Longido2us taniwha n. sp. 4. Nygolaimidac; Description of Sectomena taiaha n.sp. 5. Blon,l.iridae; Description of Dorylaimellus monticolus n. sp. 6. Loptonchide; Decription of Amphorostomum saccatum n. gen. et SD. VIII. Conclusion References Appendix I Enoplid genera recognised in New Zealand sampls ACKNOWLEDGMENTS I thank Professor B. G. Peters and Dr C. Potter for acting as my supervisors and for discussions and encourage- ment. I am grateful to the staff of the Department of Nematology, Rothamsted Experimental Station, for their kind- ness during my stay at that Institution. Especial thanks are due to Mr F. G. W. Jones for his encouragement, and to Dr J, B. Goodey for much help, advice, and free access to his excellent library. Finally, I thank the New Zealand Deoartment of Scientific and Industrial Research for financial support. NOTE References to figures) tables etc. are numbered consecutively in the type-written part, but each of the printed sections has its own series of figure numbers and so on. References to iublications follow the same scheme; all papers cited in the typewritten sections are listed in the section "References" at the end of the thesis. Papers cited in the orinted narts will be found at the end of the relevant' section. Section INTRODUCTION When this study was first considered the idea was to investigate the species composition of nematode faunas and the abundance of different species in soil and litter of a variety of New Zealand soils. Further, it was intended to investigate the effect of different vegetation regimes on the nematode fauna of the same soil types. It was also thought that it might prove instructive to examine the nema- tode faunas which have developed in the soils in the central part of the North Island of New Zealand. These soils are largely derived from showers of volcanic ash, the time of deposition of which has been recorded in recent times, or accurately determined by other workers using radio-active carbon dating techniques. These ash-showers may have caused cataclysmic destruction of the fauna when and where they occurred, and have influenced its composition since. They may also have acted as barriers to migration for a consider- able time. Lee (1959) has shown that the distribution of Megascolecine earthworms was influenced in this way. Given (pers. comm.) found a similar influence in the distribution of the Melolonthid beetles, and Forster (1954) recorded a similar discontinuity in the distribution of the harvest spiders of the suborder Laniatores. No attempt was made whilst in New Zealand to identify - 2 - any of the material collected. The collecting programme was worked out using Taylor's (1948) "Soil Map of "New Zealand" and the genetic classification of New Zealand soils included in it and in Taylor and Cox (1956). In practice the original aims proved unattainable because of the dearth of knowledge of the free-living nema- todes of New Zealand. The diversity of species encountered is illustrated by throe samples taken from Wharakohe Silt Loam (a mature podzol derived froril silicified claystones under A,athis australia Salisb. forest in a warm humid climate). The samples wore taken. under natural heath-type vegetation, from an 18-year old Pasoalum dominant pasture, and from a two-year old ryegrass--white clover pasture. Each sample yielded eighteen to twenty fairly common S10,3CiS, but only two or three species were common to any pair of samples, and none common to all. Only one species of the species examined was referable to a known species. In view of the diversity of species and the indications that the vast majority of them appeared to be n:w, and in some cases, required the erection of new genera and possibly new families, it became necessary to turn to identification and description of species rather than to continue with predominantly ecological studies as planned. Once the systematic study of the New Zealand species, and where necessary the European ones, was well in hand it - 3 - became apparent that the existing accepted arrangement of genera and species in some groups had become inadequate over the years. Much time, therefore, had to be spent incorpora- ting new information, re-examining many accepted concepts and modifying the taxonomy to fit. mild steel rubber hose 1.5 in. o.d. stainless steel tubing— -mild steel brass _tempered _ steel Fig. 1. Construction of the soil sampler. Section II MATERIALS AND METHODS 1. Materials All the New Zealand material examined was collected by the author during the period January to May 1958. Of the material from sites outside New Zealand, some was collected by the author, but much had been collected by other workers; most of it was already in the slide collection of the Department of Nematology, Rothamsted 'Experimental Station. A few specimens were borrowed from workers outside Britain. 2. Sampling Technique and Procedure As pointed out in the introduction, one of the original purposes of the study was to compare the abundance and siecios comnosition of nematode faunas under different conditions of soil type and vegetation. Therefore, an attempt was made to take sam3les of comparable size. A tubular soil sampler (fig. 1) was constructed of stainless steel because of the ease with which this material may be kept clean and the risks of cross contamination of samples reduced. The sampler was thrust into the soil to a depth of six inches, rotated, bent over at an angle to break the soil column, and withdrawn. The cutting edge was purposely made narrower than the internal diameter of the sampler body to reduce compaction of the soil core.